103

WILDLIFE BIOLOGY 9:2 (2003)

Wolf Canis lupus numbers, diet and damage to livestock in relation
to hunting and ungulate abundance in northeastern Belarus
during 1990-2000
Vadim E. Sidorovich, Larisa L. Tikhomirova & Bogumia Jedrzejewska
Sidorovich, V.E., Tikhomirova L.L. & Jedrzejewska, B. 2003: Wolf Canis lupus
numbers, diet and damage to livestock in relation to hunting and ungulate abun-
dance in northeastern Belarus during 1990-2000. - Wildl. Biol. 9: 103-111.
Wolf Canis lupus relationships with wild ungulates, domestic animals and humans
were studied in an area of ca 800 km
2
at the head of the Lovat River in north-
eastern Belarus during 1990-2000. The region was dominated by natural habi-
tats (78%) consisting mainly of forests and bogs, but also lakes and rivers. The
abundance of wild ungulates, such as moose Alces alces, wild boar Sus scro-
fa, and roe deer Capreolus capreolus, as censused by snow tracking and
assessed by game wardens, declined 5 to 6-fold between 1990 and 1996, most
probably due to uncontrolled exploitation and poaching. During 1997-2000,
the numbers of ungulates began to recover. Wolves responded to the shortage
of wild ungulates by a strong shift in feeding habits. When wild ungulates were
numerous, wolf diet as studied by scat analysis was composed of wild ungu-
lates (80-88% of consumed biomass), with small additions of medium- and small-
sized wild animals (7-13%), mainly beaver Castor fiber and hare Lepus sp.,
and domestic animals (4-6%), mainly cattle. In the years when the recorded
numbers of wild ungulates were at their lowest, wolves preyed on domestic ani-
mals (38% of biomass consumed), wild ungulates (32%), and medium- and small-
sized wild prey (29%). Wolf damage to domestic animals (28 head of cattle and
247 dogs killed) and wolf-human interaction (100 observations of wolves in
and near villages, including one attack by a rabid wolf on 11 people) were record-
ed in 14 villages. The rate of wolf predation on domestic animals and their appear-
ances in villages increased exponentially with the declining biomass of wild
ungulates and ceased again when wild ungulates began to recover; a one-year
time lag in wolf response to changes in ungulate abundance was observed. The
numbers of wolves as estimated by snow tracking and assessed by game war-
dens played a weaker role in shaping wolf-livestock and wolf-human interac-
tion. The wolf population was strongly affected by hunting during the study.
Wolves responded numerically with a 1 to 2-year time lag to the varying
intensity of harvest by humans. Our study showed the role of the human fac-
tor in shaping wolf numbers and wolf-livestock interaction in eastern Europe.
The three major components of this relationship were: 1) the manifold decline
in wild ungulate abundance, which was most probably caused by uncontrolled
exploitation by humans in the years of political transformation and econom-
ic regress, made wolves shift to predation on domestic animals; inevitably, wolves
were frequently seen in the rural areas; 2) people interpreted the growing
rates of wolf damage and appearances near the settlements as an effect of great-
ly increasing numbers of wolves, and demanded that authorities and hunters
fight the 'wolf plague'; 3) hunting impact on wolves increased and led to a marked
reduction in wolf numbers and a decline in wolf-human conflicts. This scenar-
io was most probably repeated in many areas of eastern Europe during 1990-
2000, which was a decade of political and economical transformation. From
104
WILDLIFE BIOLOGY 9:2 (2003)
The relationships of European wolves Canis lupus with
their main prey, wild ungulates, have long been affect-
ed by the events in human history and economy. First
and most important, domestic animals kept in rural
areas, from small poultry to horses, offered an impor-
tant alternative prey to wolves, the more so as wild un-
gulates declined in range and abundance. Secondly,
hunting harvest of both ungulates and wolves affects wolf
numerical responses to variation in ungulate abundance.
Meriggi & Lovari (1996), having analysed data on wolf
diets in 15 localities in southern Europe, showed that the
shares of wild and domestic ungulates in wolf diets were
strongly negatively correlated. Indeed, in the whole
continent, species composition of wolf prey varied
from almost exclusively wild ungulates belonging to five
native species in the Biaowiea Primeval Forest, east-
ern Poland (Je drzejewski, Je drzejewska, Okarma,
Schmidt, Zub & Musiani 2000) to domestic ungulates
of five species in Greece (Papageorgiou, Vlachos, Sfouga-
ris & Tsachalidis 1994). Meriggi, Brangi, Matteucci &
Sacchi (1996) documented that, in Italy, the use of
livestock species was lower wherever the richness and
diversity of wild ungulate guild increased. The same pat-
tern, but in the temporal scale, was described by Jedrze-
jewska & Je drzejewski (1998) from central-eastern
Europe. As pasturing of cattle in the woodland ceased
and numbers of wild ungulates recovered, wolves eas-
ily shifted from frequent hunting on livestock to prey-
ing on wild ungulates only. Ryabov (1974) reported
that, in the Voronezh region in Russia, wolves caused
little damage to numerous goats and other livestock,
because free-living sika deer Cervus nippon (introduc-
ed to the Khopersk Nature Reserve) were very abundant.
Livestock predation by wolves may also be affected
by human-imposed changes in the abundance of wild
ungulates. Jedrzejewska, Jedrzejewski, Bunevich, Mi-
kowski & Krasi nski (1997) analysed the population
dynamics of five species of wild ungulates in the Bia-
owiea Primeval Forest in eastern Poland and western
Belarus based on data covering >100 years and found
that during periods of war, military and political unrest
and economical regress, wild ungulates served as food
resources for humans and suffered heavy poaching that
could even lead to extermination of the largest, and
most slowly reproducing species, e.g. European bison
Bison bonasus. In such situations, wolves may have to
compensate for the decline in wild prey by hunting
domestic ungulates where and whenever available (Os-
molovskaya & Priklonskii 1975).
In this paper, we report 11 years of monitoring data
on the relationship between wolves, wild ungulates
and livestock in natural woodlands interspersed with ru-
ral areas in northeastern Belarus. The study period
(1990-2000) was characterised by profound political
changes and economical regress. This situation favoured
uncontrolled exploitation of wild ungulates, which led
to their fast, many-fold decline. In this paper, we analyse:
1) dietary response of wolves to changes in wild ungu-
late abundance, 2) effect of harvest by humans on wolf
numbers, and 3) frequency of wolf attacks on domes-
tic animals.
a management perspective, we suggested that predation levels and wolf-human
conflicts could be reduced not only by increased wolf harvest but also by enhanc-
ing the density and diversity of wild ungulates.
Key words: Canis lupus, feeding habits, livestock depredation, moose, roe deer,
wild boar, wolf-human interaction
Vadim E. Sidorovich & Larisa L. Tikhomirova, The Vertebrate Predation
Research Group, Institute of Zoology, National Academy of Sciences of Belarus,
Akademicheskaya Str. 27, 220072 Minsk, Belarus - e-mail address: vadimsid@
mustbel.open.by
Bogumi a Je drzejewska, Mammal Research Institute, Polish Academy of
Sciences, 17-230 Bia owiea, Poland - e-mail: bjedrzej@bison.zbs.bialowieza.pl
Corresponding author: Vadim E. Sidorovich
Received 10 October 2001, accepted 20 September 2002
Associate Editor: Henryk Okarma
105
WILDLIFE BIOLOGY 9:2 (2003)
Study area
The study was carried out in an area of about 800 km
2
at the head of the Lovat River (5545'N, 3020'E) in the
Gorodok district, Vitebsk region, northeastern Belarus
(Fig. 1). The study area is characterised by large forests
and bogs on rough glacial terrain, fairly natural network
of rivers and glacial lakes. Natural habitats comprise
about 78% of the area, and man-transformed land (i.e.
fields, pastures, meadows, human settlements and roads)
covers 22%, in some localities up to 31%. Agricultural
land is dominated by dry meadows (16% of the entire
region, and 73% of the agricultural land). The average
density of the human population is about 3 people/km
2
.
The study area belongs to the transitional region of
temperate mixed forests. Spruce Picea abies and pine
Pinus sylvestris are the dominant species among the
coniferous trees. Alder Alnus glutinosa and A. incana,
birch Betula pendula and B. pubescens, and aspen
Populus tremula are the most common deciduous trees.
Masting deciduous trees such as oak Quercus robur and
lime Tilia cordata are rare. The guild of large predators
includes wolf, lynx Lynx lynx, and brown bear Ursus arc-
tos. The community of wild ungulates consists of three
species: moose Alces alces, roe deer Capreolus capre-
olus, and wild boar Sus scrofa. Other possible prey
species of wolves were beavers Castor fiber, blue hare
Lepus timidus, and brown hare L. europaeus.
The terrain is flat to undulating with altitudes lying
within 100-200 m a.s.l. Mean temperature in July is
18C, and in January -6C. Winters are variable, but usu-
ally snow cover persists for at least 1.5-2 months, with
a maximal depth of 30-90 cm. Maximum duration of the
snow period is about six months, i.e. from late October
till mid April. During winter, periods of severe frost
(below -20C) alternate with thaws lasting for some
weeks.
Material and methods
In 1990-2000, abundance of wolves and wild ungulates
was estimated by snow tracking. Each year in late win-
ter (January-March), the same 36-km transect, located
in the natural habitats, was walked, and the number of
animal trails crossing the transect was counted. The route
crossed various forest types, river val-
ley, lake shore and open marshes as
well as rural areas. It was represen-
tative of the landscape structure and
composition in the whole study area.
The abundance index of each species
was calculated as the number of tracks
divided by the number of kilometres
walked and the number of days passed
from the last snowfall, and multiplied
by 2 (number of tracks/ km/2 days).
As an additional source of data, esti-
mates of wolf and ungulate numbers
from the local department of the Be-
larusian hunting organisation (Gorodok
Raisovet BOOR) were also used for
analysis. The estimates come from
game wardens and forest guards, who
inspect their districts many times dur-
ing each winter (e.g. before and dur-
ing hunts for ungulates and wolves,
when hunting for furbearers, and when
inspecting the signs of poaching activ-
ities) and - by recording tracks, visu-
al observations, and other signs of
animal activities - estimate the number
of ungulates and wolves in their dis-
tricts (covering a total of 728 km
2
).
Number of wolves and ungulates ob-
Figure 1. The study area is located in the basin of the Lovat River in the Gorodok district, Vitebsk
region, northeastern Belarus. Names of villages are those from which data on wolf sightings
and predation on domestic animals were collected.
0 2 4 6 8 10 km
30 15
o
5
5
5
0

o
Sennitsa
Lake
Prudok
Bolotnitsa
Dorokhi
Zadrache
Bibino
Nemtsevo
Rudnia
Borki
Kaverzy
Lyakhovka
Makhalovo
Sharipy
Mezha
R
U
S
S
I
A
B
E
L
A
R
U
S
BY
RU
Forests and marshes
Lakes
Rivers
Roads
Villages
State border
106
WILDLIFE BIOLOGY 9:2 (2003)
tained from the hunters organisation was recalculated
per 100 km
2
. Based on personal experience of game war-
dens rather than on methodical surveys, this estimate is
an index of abundance and not the actual density. We
believe, nonetheless, that it well reflects the multiannual
trend in numbers as the data were gathered in the same
way throughout the study period. For calculation of crude
biomass of wild ungulates the following body masses
were applied: for moose 200 kg, for wild boar 80 kg,
and for roe deer 20 kg.
Wolf predation on livestock and wolf sightings by
humans around villages were monitored in an area of
about 240 km
2
located at the centre of the whole study
region, and populated by humans more densely than the
other parts of it. Large patches of natural habitats were
interspersed with the 14 villages Bibino, Bolotnitsa, Bor-
ki, Dorokhi, Kaverzy, Lyakhovka, Makhalovo, Mezha,
Morozovo, Nemtsevo, Prudok, Rudnia, Sharipy and
Zadrache (see Fig. 1). The majority of these villages were
small (5-26 houses and 9-67 inhabitants), but two were
fairly large (49 and 84 houses, 147 and 242 inhabitants).
Farmers mostly kept cows, pigs, hens, dogs and cats,
whereas horses, sheep and geese were less common. Two
collective farms ('kolkhoz') operated in the two large vil-
lages, and they had many cows (110 and 280), pigs (50
and 120), and horses (15 and 40). In each village, we
contacted an experienced and reliable person, who
gathered information about cases of wolf damage on live-
stock and other observations on wolves in the village
and its surroundings. A total of 377 such observations
were collected during 1990-2000.
In order to analyse the diet composition of wolves, their
scats were collected in the study area, mainly along for-
est paths. During 1990-1992, 1994-1996 and 1999-
2000, a total of 1,185 scats were gathered and ana-
lysed. Almost equal numbers of scats came from spring-
summer and autumn-winter seasons. Analysis of scats
followed the standard procedure (Lockie 1959, Gosz-
czy nski 1974, Je drzejewska & Je drzejewski 1998);
they were washed through a 0.5-mm mesh sieve and the
remaining contents were identified to species based on
bone and tooth fragments (according to Pucek 1981) and
microscopically, using the keys for mammalian hair
(Debrot, Mermod, Fivaz & Weber 1982, Teerink 1991).
The relative amounts of the various prey species in
wolf diet were given as percentage of occurrence in the
total number of analysed scats (%Occ) and percentage
of biomass consumed (%Bio) by wolves. To assess the
consumed biomass of each species, the weight of prey
remains recovered from faecal samples was multiplied
by coefficients of digestibility. Since no such digestibil-
ity factors for wolves are available, data for the relat-
ed canid species, the red fox Vulpes vulpes, were ap-
plied (after Goszczy nski 1974, Jedrzejewska & Jedrze-
jewski 1998). Food niche breadth (B index; after Levins
1968) was calculated for the percentage of biomass con-
sumed in five food categories:
B = 1/p
i
2
,
where p
i
is the fraction of a given food category in the
total prey biomass consumed by wolves. B could vary
from 1 (the narrowest niche) to 5 (the broadest niche pos-
sible). To assess wolf preferences for the three species
of wild ungulates, Ivlevs selectivity index D (after
Jacobs 1974) was calculated:
D = (r - p)/(r + p - 2rp),
where r is the fraction of the species among wild ungu-
lates consumed by wolves (total number of findings of
wild ungulate remains in all scats was taken as 100 to
calculate the fractions), and p is the fraction of the spe-
cies in the wild ungulate community (calculated from
pooled data of track counts). D can range from -1 (total
avoidance of a species) to 0 (selection proportional to
occurrence) to 1 (maximum positive selection).
Results
Population trends of wild ungulates
Estimates of ungulate abundance in 1990-2000 obtained
by the two methods (snowtracking on transects and as-
sessments by game wardens) were strongly mutually cor-
related (moose: r = 0.96, N = 11 years, P < 0.0005; roe
deer: r = 0.94, P < 0.005; wild boar: r = 0.97, P <
0.0005), suggesting that they reflected the population
trends of the three species. Moose steadily declined with
the minimum numbers, in 1994-1996, being one-tenth
of those recorded in 1990 (Table 1). After 1997, moose
numbers grew, and in 2000 they reached 40-50% of the
initial numbers. In contrast, roe deer densities were
low during 1990-1995, and then they increased 5 to 10-
fold until 2000 (see Table 1). The wild boar, always a
dominating species in the wild ungulate community,
declined >10-fold from 1990 to 1996 and recovered in
1997-2000 to reach 60-80% of their initial densities (see
Table 1). The population of each species showed different
dynamics. If pooled together, the numbers of all ungu-
lates declined from 1990, were at their lowest level in
1994-1996, and started to increase after that time (Fig.
2). During the 11 years of our study, the amplitude of
variation was very large: 4 to 5-fold in estimated num-
107
WILDLIFE BIOLOGY 9:2 (2003)
bers of wild ungulates, and 7-fold in their crude biomass.
The species structure of the community also varied
during the study period. The share of moose in the
total numbers of wild ungulates declined from 20% in
1990 to 13% in 1996 and 9% in 2000. The proportion
of roe deer grew from 4% in 1990 to 50-53% in 1996-
2000, and the share of wild boar declined from 76% in
1990 to 34% in 1996, and then increased to 41% in 2000.
However, in terms of biomass, the wild boar dominat-
ed throughout the study period (percentage shares in
1990, 1996 and 2000 were 60, 43 and 55%, respectively),
followed by moose (39, 40 and 29%, respectively).
Roe deer contributed the least to the total crude biomass
of wild ungulates (1, 17 and 16% in 1990, 1996 and
2000, respectively).
Table 1. Index of wild ungulate winter abundance (tracks/1-km transect/2 days), index of estimated ungulate winter abundance (individu-
als/100 km
2
), index of wolf winter abundance and hunting harvest, and wolf interaction with livestock and humans in the head region of
the Lovat River (Gorodok district, Vitebsk region, NE Belarus) during 1990-2000.
Year
Species/records 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000
Index of ungulate winter abundance
Moose 12.7 10.1 7.6 2.1 0.9 1.3 1.2 2.8 3.7 4.5 7.1
Roe deer 1.3 0.9 1.1 1.0 1.5 1.1 2.9 8.2 10.6 16.2 13.6
Wild boar 25.1 27.4 18.1 10.1 4.2 2.1 1.5 6.1 10.0 18.1 20.9
Estimate of ungulate winter abundance
Moose 61 52 34 16 7 7 9 20 17 28 23
Roe deer 12 20 18 24 36 32 38 54 96 124 132
Wild boar 238 201 154 98 51 48 24 56 96 142 110
Index of wolf winter abundance and wolf harvest
Wolf tracks
a
1.4 0.6 1.0 0.3 0.7 1.1 2.3 1.6 0.5 0.4 0.6
Wolf numbers
b
2.2 1.3 1.5 0.9 1.6 3.6 2.1 3.0 0.5 0.8 1.7
Wolves shot
c
24 20 28 7 9 5 27 19 15 10 12
Wolf interaction with livestock and humans
Wolves seen in village 1 2 4 0 2 5 29 37 12 3 5
Cattle killed by wolf 0 0 1 0 2 5 12 4 1 2 1
Dogs killed by wolves 2 1 2 6 24 44 72 55 21 13 7
a
Number of tracks/1-km transect/2 days;
b
Number of wolves/100 km
2
based on game wardens assessment of the numbers of wolves in winter;
c
Number of wolves shot by hunters in the Gorodok district.
N
U
M
B
E
R
80
70
60
50
40
30
20
10
0
1990 91 92 93 94 95 96 97 98 99 2000
Dogs killed by wolves
Observations of wolves
in the village areas
Cattle killed by wolves
A)
40
30
20
10
0
400
300
200
100
0
1990 91 92 93 94 95 96 97 98 99 2000
Density of wild ungulates
Biomass of wild ungulates
N
U
M
B
E
R

O
F

W
I
L
D

U
N
G
U
L
A
T
E
S
C
R
U
D
E

B
I
O
M
A
S
S

O
F

W
I
L
D

U
N
G
U
L
A
T
E
S
YEAR
B)
Figure 2. Number of dogs (M) and cattle (I) killed by wolves, and obser-
vations of wolves in and near villages (; A) in relation to the abun-
dance expressed as density (G; number of individuals/100 km
2
) and crude
biomass (G; in tonnes/100 km
2
) of wild ungulates (B) pooled for
moose, roe deer and wild boar, and censused in an area of 728 km
2
.
Wolf-livestock and wolf-human interaction were monitored in an area
of 240 km
2
embracing the 14 villages (data as in Table 1).
Figure 3. Numerical response of wolves expressed as mean number of
wolf tracks/km/2 days in relation to the mean number of wolves shot
by hunters in the region (with a 1 to 2-year lag) in 1990-2000 (data as
in Table 1). The arrows show the time sequence of events.
3.0
2.5
2.0
1.5
1.0
0.5
0
0 5 10 15 20 25 30
NUMBER OF WOLVES SHOT BY HUNTERS
M
E
A
N

N
U
M
B
E
R

O
F

W
O
L
F

T
R
A
C
K
S
/
K
M
/
2

D
A
Y
S

108
WILDLIFE BIOLOGY 9:2 (2003)
Wolf response to hunting harvest by humans
and abundance of wild ungulates
Estimates of wolf numbers obtained by snowtracking
and game wardens assessments were mutually corre-
lated (r = 0.63, N = 11 years, P = 0.04). The mean esti-
mate of wolf density during the study was 1.75 indi-
viduals/100 km
2
(SD: 0.94). Changes in wolf numbers
were not correlated with the variation in wild ungulate
densities, neither directly nor with 1 or 2-year time
lags (P > 0.1). The most important factor shaping the
variation in wolf numbers was the mean hunting har-
vest of wolves by humans in the preceding two years
(R
2
= 0.43, N = 9 years, P = 0.05 for wolf tracks, and
R
2
= 0.50, P = 0.03 for estimated wolf densities; see data
in Table 1). Thus, wolves responded to the changing rate
of persecution with a 1 to 2-year time lag (Fig. 3).
However, variation in wild ungulate numbers did
have a strong impact on prey searching and hunting
behaviour of wolves. As wild ungulates declined, wolves
were more likely to approach the vicinities of human set-
tlements in search for domestic animals (see Fig. 2).
During 1990-2000, 28 cows and 247 dogs were report-
ed to have been killed by wolves. Among the dogs, 117
(47%) were hunters dogs attacked by wolves when
roaming in the field or forest away from the village, and
130 (53%) were watch dogs, often chained, killed by
wolves near the houses. Moreover, in 1990-2000, wolves
were observed 100 times in the village areas. These
observations include the case of a rabid wolf that, in
1997, came into one of the villages and bit 11 people
before it was killed by locals. The only other possibly
direct interaction of wolves and humans happened in
1995, when a dead man was found in the forest, his body
bearing signs of having been scavenged presumably by
wolves. As the man had been known to suffer from heart
disease, the ultimate cause of his death was unclear.
Table 2. Roles of ungulate abundance (with 1-year lag) and wolf numbers in shaping wolf interaction with domestic animals and humans.
The results of the multiple regression analyses are based on data from Table 1; N = 10 years, *** P < 0.001, ** P < 0.01, * P < 0.05, ns -
not significant. Semipartial correlations squared (sr
i
2
) show the contribution of each dependent variable to the total variation explained (Tabachnick
& Fidell 1983).
Contributions of independent variables to the total variation explained (sr
i
2
)
Dependent variable Total variation Biomass of wild
log-transformed, ln(x+1) explained R
2
ungulates in year n-1 Index of wolf abundance in year n
Number of dogs killed 0.991*** 0.587*** 0.020** (Tracks)
0.976*** 0.695*** 0.007 (ns) (Numbers)
Number of cattle killed 0.896*** 0.180** 0.274** (Tracks)
0.713** 0.310* 0.091 (ns) (Numbers)
Number of observations of 0.682** 0.069 (ns) 0.301* (Tracks)
wolves in the village area 0.412 (ns) 0.215 (ns) 0.031 (ns) (Numbers)
Table 3. Diet composition of wolves in the periods of high (1990-1992 and 1999-2000) and low (1994-1996) abundance of wild ungulates
expressed as percentage occurrence in scats (%Occ), and percentage of consumed biomass (%Bio). + = contribution of <0.05%. Food niche
breadth B (after Levins 1968) calculated for four categories of food: wild ungulates, wild medium and small-sized prey, domestic medium
and small-sized animals and plant material.
1990-1992 1994-1996 1999-2000
Prey category %Occ %Bio %Occ %Bio %Occ %Bio
Moose Alces alces 42.7 32.5 19.8 10.8 14.9 9.7
Roe deer Capreolus capreolus 6.5 3.6 10.5 4.0 20.3 10.2
Wild boar Sus scrofa 87.7 52.3 26.2 17.3 85.3 60.1
Total wild ungulates 96.1 88.4 54.0 32.1 93.3 80.0
Raccoon dog Nyctereutes procyonoides 2.9 0.9 25.1 5.9 15.5 3.2
Hare Lepus sp. 5.8 2.2 53.4 17.2 18.9 4.7
Beaver Castor fiber 6.3 3.3 22.6 5.1 10.7 4.8
Micromammalia 1.1 + 10.5 0.3 1.9 +
Wild birds Aves 4.7 0.9 6.5 1.0 3.5 0.3
Total wild medium and small-sized prey 17.9 7.3 70.5 29.5 36.8 13.0
Domestic cattle 6.3 3.4 35.8 23.6 8.0 4.7
Other domestic ungulates 0.6 0.3 5.0 3.1 1.1 0.8
Domestic dog Canis familiaris 0.9 0.3 21.2 9.1 2.9 0.9
Domestic cat Felis catus 0.2 + 10.7 1.9 0.8 0.1
Domestic birds - - 0.8 0.1 - -
Total domestic animals 8.1 4.0 74.9 37.8 12.8 6.5
Fruits and seeds 1.6 0.2 2.8 0.5 2.9 0.4
Other plant material 1.1 0.1 2.5 0.1 1.3 0.1
Food niche breadth B 1.27 3.65 1.52
Mean number of prey/scat 1.69 2.37 1.88
Number of scats analysed 447 363 375
109
WILDLIFE BIOLOGY 9:2 (2003)
The number of cattle and domestic dogs killed by
wolves, as well as observations of wolves around the vil-
lages increased exponentially with the decline in wild
ungulates and the highest number of such incidences was
recorded during 1995-1997. Indeed, the between-year
variation in wolf interaction with domestic animals
and humans were best explained by two factors: abun-
dance (biomass) of wild ungulates with a 1-year time
lag and index of wolf abundance (multiple regression
analyses: R
2
from 0.991 to 0.412, P from <0.0005 to 0.15,
N = 10 years). Semipartial correlations squared calcu-
lated for each regression model showed that the abun-
dance of wild ungulates (acting with a 1-year lag) was
a far more important factor than wolf numbers (Table
2).
The diet composition of wolves also reflected the fall
and rise of wild ungulates in the study period. Wild ungu-
lates constituted 88% of food biomass consumed by
wolves during 1990-1992, 32% in 1994-1996, and 80%
during 1999-2000 (Table 3). The most important buffer
prey compensating for the shortage of wild ungulates
was domestic animals (mainly cattle) and wild medium-
sized mammals (predominantly hare, but also raccoon
dog Nyctereutes procyonoides and beaver).
Wolves preyed on the three wild ungulates oppor-
tunistically, with no consistent preferences for any spe-
cies. Their percentage shares in the total food biomass
consumed by wolves were proportional to the amount
of crude biomass of wild ungulates available (Fig. 4A).
Also, the species structure of ungulates preyed upon
reflected that in the living community (Fig. 4B). Food
niche breadth calculated for five groups of prey (see Table
3) was the narrowest in 1990-1992, when wild ungu-
lates were most numerous (B = 1.27) and the widest in
1994-1996 (B = 3.65), when they were rare.
Discussion
In Europe, the numerical response of wolves to varia-
tion in ungulate abundance is even more disturbed by
humans than their dietary, functional response. Until
very recently, deliberate control or eradication of
wolves has been an indisputable, widely accepted way
of wolf 'management'. The only periods when wolves
could recover in numbers and regain some of their
original range were the times of war and uprisings
(Osmolovskaya & Priklonskii 1975, Je drzejewska,
Jedrzejewski, Bunevich, Mikowski & Okarma 1996).
In effect, in many regions of central and eastern Europe
the negative correlation between wolf numbers and
ungulate densities was observed (Filonov 1989, Jedrze-
jewska et al. 1997) instead of a typical positive numer-
ical response. In our study, both wolf and ungulate
dynamics were shaped predominantly by human har-
vesting. Though we have no precise data on ungulate
hunting quotas and poaching intensity, the information
obtained by interviewing the local hunters as well as
our observations on the remnants of the poached moose
and wild boar in the study area allowed us to roughly
estimate that, since 1992, the killing rate of ungulates
by humans could have increased 5 to 10-fold in com-
Figure 4. Shares of the ungulate species moose (I), roe deer (M) and
wild boar (G) in the total biomass consumed by wolves (in %) in rela-
tion to the crude biomass of the respective species (A; in tonnes/100
km
2
) and as their percentage share of the total biomass of all ungulates
(B). The three points for each species denote mean values for the peri-
ods 1990-1992, 1994-1996 and 1999-2000, respectively. For A): y =
2.64 + 3.59x; R
2
= 0.811. P = 0.001, and for B): y = 1.90 + 1.06x, R
2
=
0.865, P < 0.0005.
Wild boar Moose Roe deer
100
80
60
40
20
0
0 5 10 15 20
S
H
A
R
E

O
F

U
N
G
U
L
A
T
E

S
P
E
C
I
E
S

O
F

T
O
T
A
L

B
I
O
M
A
S
S

C
O
N
S
U
M
E
D

B
Y

W
O
L
V
E
S
CRUDE BIOMASS
%
A)
0 20 40 60 80 100
100
80
60
40
20
0
S
H
A
R
E

O
F

U
N
G
U
L
A
T
E

S
P
E
C
I
E
S

O
F

T
O
T
A
L
B
I
O
M
A
S
S

C
O
N
S
U
M
E
D

B
Y

W
O
L
V
E
S
PERCENTAGE SHARE
%
B)
110
WILDLIFE BIOLOGY 9:2 (2003)
parison with earlier years and it remained heavy until
1995. Thus, uncontrolled exploitation by humans was
the most probable cause of the ungulate decline in the
1990s. Conducive to this was the political transforma-
tion and economical regress in Belarus (before 1991 -
the Belarusian Socialist Soviet Republic).
Our study evidenced that a manifold decline in wild
ungulate abundance was an essential factor responsible
for the increased predation on domestic animals by
wolves. Interestingly, however, our interviews with
local people revealed that they interpreted the observed
changes as resulting from presumed growth of the wolf
population. In 1990-1993, when wolves only rarely
killed cattle or dogs, local people forgot about wolf dan-
ger to livestock and even started to think that wolves must
have been rare in the area. When wolves began to ap-
pear in and near the villages, the local citizens believed
that this was the result of a marked increase in wolf num-
bers. Such interpretation of wolf-livestock interaction
was a common opinion in many areas of Belarus in the
decade 1990-2000, and was the main reason for inten-
sified hunting of wolves after 1996.
The number of wolves did not decline with reduced
abundance of wild ungulates, because the wolves had
access to numerous domestic animals. In the early
1990s, wolves were relieved from heavy control but few
years later, when heavy damage to livestock caused
complaints from farmers, hunting on wolves intensified.
Worth attention is the shape of the wolfs numerical
response to hunting by man. It occurred with a substantial
(1-2 years) time lag as could be expected from the re-
production and maturation rate of wolves. Also, a one-
year lag was observed in the dietary shifts of wolves to
domestic animals when wild ungulates declined. The
wolves continued to prey on declining wild ungulates,
but then again, when wild ungulates began to recover,
wolves for some time continued their rallies to the vil-
lages before they shifted back to wild prey. This sug-
gests substantial conservationism in wolf hunting habits.
The described situation of ungulate-human-wolf rela-
tionship seems typical for the large regions of Belarus
in the 1990s and most probably occurred in many areas
of eastern Europe during 1990-2000, as this was a
decade of profound political and economical transfor-
mation. From a management perspective, we suggest that
predation level and human-wolf conflicts could be re-
duced not only by increased wolf harvest in agricultu-
ral areas (Mech 2001), but also by enhancing the den-
sity and diversity of wild ungulates. A similar conclu-
sion was drawn in a study on Portugese wolves (Vos
2000) which fed exclusively on livestock because of the
scarcity of wild prey. Vos (2000) emphasised that rein-
troduction of native prey species is essential for con-
servation of wolves in Portugal.
Finally, the controversial issue of wolf attacks on
humans still awaits serious analysis and explanation.
Osmolovskaya & Priklonskii (1975) and Ryabov (1988)
proposed that such incidences were more likely to hap-
pen, when wolves became numerous and suffered dead-
ly shortage of food, when rabies spread in the wolf pop-
ulation, and when hybridisation with stray dogs was
common.
Acknowledgements - we are grateful to Vyacheslaw Budny,
Dmitrii Petrov, Genadii Lauzhel and Aleksei Polozov for their
help in the field work, to Karol Zub for drawing the figures,
and to Dr Wodzimierz Jedrzejewski for his comments on an
earlier draft of this paper. The study was financed by the
Institute of Zoology, National Academy of Sciences of Belarus.
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